Figure 1. The Nesbitt (2011) tree with the Archosauria highlighted in yellow and Lewisuchus in blue. Taxa not in the Archosauria, according to the large reptile tree, are highlighted in orange. The large reptile tree found a similar nesting for Lewisuchus, but with the addition of more taxa, more resolution nests just crocs (including early bipeds like Scleromochlus) with dinos (which also includes silesaurs and poposaurs. Rauisuchians nest further back, closer to Euparkeria and erythrosuchids.

In the Nesbitt (2011) tree (Fig. 1) the Archosauria also includes the rauisuchians, pterosaurs, lagerpetids, ornithosuchians, aetosaurs, poposaurs (allied with Qianosuchus and the basal rauisuchian, Arizonasaurus). I thought this may have been part of the problem (see below). Furthermore, Nesbitt (2011) did not include some of the basal crocs, like Pseudhesperosuchus, Decuriasuchusand Scleromochlus. He also did not include Vjushkovia, a basal rauisuchian close to the basal crocs and basal dinosaurs. Without these key basal taxa the derived taxa are further from one another, making it more difficult to assess the gradual accumulation of traits we’re all looking for.

So, why not eliminate some of the strange bedfellows?
If we remove Mesosuchus, the Pararchosauriforms, Vancleavea and pterosaurs, we’re left with pretty much the same tree (Fig. 2). This pruned tree is in closer agreement with the large reptile tree except at the base of the Archosauria, where the aforementioned missing taxa would have nested and perhaps shifted things around a bit. Who knows? As noted earlier, the addition of several Youngina brings resolution to this problem, but it was excluded by Nesbitt (2011).

CM73372
Since CM73372 appears at the base of the crocs, just beyond the standard rauisuchia, and was first labeled a juvenile Postosuchus, I’m keenly interested in seeing this, but so far have not. Requests have been sent. No replies yet. Any jpegs would be welcome.

Figure 2. The Nesbitt tree without Mesosuchus, pterosaurs, pararchosauriformes and Vancleavea, taxa that belong elsewhere according to the large reptile tree. It holds together pretty well. Here Marasuchus arises more or less directly from Euparkeria. Here Lewisuchus is not so far from Gracilisuchus and Turfanosuchus, matching the large reptile tree results, almost.

Bipeds Galore!
According to Nesbitt (2011) the origin of the dinosaurs is led by one one biped after another (not to mention the pterosaurs, which most paleontologists ironically refuse to grant basal bipedal status to, due to Late Jurassic and Cretaceous footprints of secondarily quadrupedal beachcombers).

Figure 3. Basal bipedal dinosauriformes, from Lagerpeton through Marasuchus, Lewisuchus, Asilisaurus, Sacisaurus and Silesaurus, according to Nesbitt (2011). It’s easy to see why what little is known of Lagerpeton was lumped with these taxa (much of it due to hopeful glee), but its feet and ankles give it away as a Tropidosuchus sister.

The Large Reptile Tree Recovered Another Solution
This image was first published 11 months ago on a pterosaur heresies blog about the origin of the Dinosauria. It’s still pretty fresh. Note the gradual accumulation of traits in sisters that look more like each other than any competing set of candidates.

Figure 4. The heretical model of dinosaur origins (Peters 2007). Here basal rauisuchians gave rise to smaller bipedal crocs and dinos, which later diversified. There’s Lewisuchus at #4, just in front of tiny Scleromochlus.

There’s more about the origin of dinosaurs here. Seems Lewisuchus was not far from the origin of the Dinosauria, and may be the best candidate, but it also may be a deadend. More bipedal crocs will help us figure this out.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Nesbitt SJ 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352: 292 pp.

Tijubina pontei (Bonfim and Marques 1997) was a tiny Early Cretaceous lizard from the Crato Formation (late Aptian) of northeast Brazil. In a recent redescription Simões (2012) reported Tijubina lacked the posteroventral and posterodorsal processes of the dentary and the tibial/fibular length equaled the femoral length. Its posterior dentary teeth were robust, cylindrically based, unsculptured and bore no cuspids. Simoes (2012) nested Tijubina in a basal position among the Squamata. Reynoso (1998) reported a similar nesting for Huehuecuetzpalli. Neither considered the possibility that both specimens nested in a third squamate clade, the Tritosauria, outside of the Iguania + Scleroglossa.

Late Survivors
Both Huehuecuetzpalli and Tijubina were late survivors of a 130 million year earlier Late Permian radiation of lizards. Tijubina is distinguished by its teeth, which are larger posteriorly and shaped like cylinders instead of sharp points. Tijubina was about half the size of its Huehuecuetzpalli.

Figure 2. Tijubina in situ, nearly full size on a 72dpi screen. Click to enlarge.

Not a Juvenile
Simoes (2012) described Tijubina as immature due to a imcompletely calcified joints, a wide open sternal fontanelle (hole), unfused pectoral and pelvic elements. Adult tritosaur lizard sisters are likewise incompletely calcified. Unlike Huehuecuetzpalli, and despite its smaller size, the carpal elements of Tijubina were well ossified. The lack of dorsal and ventral processes of the posterior dentary are traits shared with Huehuecuetzpalli.

Figure 2. Manus of Tijubina identifying carpal elements. Metacarpal 4 is largely beneath mc5. Here the two centrale are ossified along with the other carpal elements and present. The carpus is unossified in adult Huehuecuetzpalli.

The carpus is not ossified in Huehuecuetzpalli, but it is well ossified in the much smaller Tijubina and both centrale are present. Earlier I wondered if the pteroid and preaxial carpal were migrating at the evolutionary stage represented by Huehuecuetzpalli because the carpus was poorly ossified. That would have been an ideal time to do it! Here Tijubina may have been a sister Huehuecuetzpalli, but the latter was closer to fenestrasaurs including pterosaurs.

Figure 3. The pelvis and possible prepubis and Tijubina. Is this the origin of the prepubis? Or just a splinter or two of bone in the position of the prepubis. I can’t tell for sure. Phylogenetically Tijubina was scored without a prepubis and pteroid.

Cosesaurusthrough pterosaurs all have a prepubis, a new bone extending beyond the ventral margin of the pubis. So the prepubis appeared some time prior to Cosesaurus. It may or may not be present in Langobardisaurus. It is not present in Huehuecuetzpalli. A possible prepubis may be present in Tijubina (Fig. 3). On the other hand, that little fleck of bone(s) may just be a splinter from the damaged pubis. No problem either way.

A Long Tibia
Since the tibia was subequal to the femur, Tijubina was likely a sprinter and a possible occasional biped, like many living lizards with similar proportions. Such traits and behaviors likely led to the development of a prepubis in sister taxa.

Figure 4. Pes of Tijubina. PILs added.

Pes
The pes of Tijubina had tendril-like toes, indicating an arboreal lifestyle. Like Huehuecuetzpalli and Cosesaurus the proximal phalanges of digit 5 were long. The tarsals were not coossified, a trait typical of many (but not all) tritosaurs. Fenestrasaurs (including pterosaurs) did not ossify two distal tarsals. Drepanosaurs and all living lizards co-ossified the proximal tarsals.

SummaryTijubina was a late-surviving representative of the Tritosauria, a clade of lizards that ultimately gave rise to tanystropheids, drepanosaurs and pterosaurs. The cylindrical teeth were autapomorphies not found in other clade members. The tiny size and crushed nature of the specimen prevent confirmation of several possible fenestrasaur-like traits.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

The smallest known pterosaurB St 1967 I 276 (No. 6 of Wellnhofer 1970 ) was discussed earlier. Today we get to meet maybe the second smallest pterosaur, Pterodactylus meyeri BMNH 42736 (Munster 1842, Fig. 1) is the same size as No. 6, but had several distinct traits (Fig. 2). I ran across the BMNH specimen in Unwin’s (2006) The Pterosaurs From Deep Time book on page 151. Dr. Unwin considered the specimen a “flapling” (= newly hatched pterosaur able to fly) with a wingspan of 17 cm, so that is the reconstructed scale (Fig. 3).

The Value of a Reconstruction
It’s a shame that modern workers don’t produce reconstructions of crushed pterosaurs anymore. There is so much to see (Figs. 2, 3), especially when one compares similar specimens. Many traits would go unnoticed if left crushed.

Figure 1. One of the world's smallest pterosaurs, traced from Unwin (2006, p. 151). The feet of the "flapling" were not articulated and a certain amount of guesswork was applied to the idenfication of the pedal elements and their reconstruction. Note how the left radius and ulna are parallel to and beneath the elongated right scapula. The right coracoid is largely beneath the right humerus. The left hand and proximal wing finger are beneath the right hand. Soft tissue stains are highlighted in orange. The wing had a narrow chord at the elbow. Colorizing the bones is a result of employing DGS, digital graphic segregation.

Distinct from No. 6, the “flapling” had a deeper skull, more and smaller dorsal vertebrae and ribs, a longer scapula (almost touched the pelvis), a deeper and more fully fused pelvis and a larger sternal complex than either of its sisters. Considering the reconstructed differences in quadrate elevation, jugal shape and pelvis dimensions (Fig. 2), you might think the “flapling” would have nested further apart from No. 6 and No. 12. These distinctions suggest that the “flapling” may have been at the base of its own clade of currently unknown descendants.

Figure 2. The tiniest pterosaurs. On the left, Unwin's "flapling" Pterodactylus meyeri BMNH 42736. On the right, B St 1967 I 276, No. 6, the former sole owner of the title.

Juvie or Adult?
If the BMNH tiny pterosaur was indeed a juvenile of a larger more established taxon, which one did it match up to? And if so, why did it nest with other tiny pterosaurs? No. The BMNH specimen was an adult. The many autapomorphies (= differences) in the “flapling” also follow a larger trend seen in other tiny pterosaurs: morphological innovation.

Figure 3. Full scale image of ginkgo leaf and the two smallest pterosaurs to scale on a 72 dpi screen. Yes, these are tiny, but just look at the size of a hatchling on the far right, no bigger than a small fly.

Special Premaxillary Teeth
In the BMNH “flapling” we see more substantial anteriorly-directed medial teeth forming the tip of the premaxilla. Those two teeth evolve to become one in the rostral tip of Germanodactylus. That tooth is the only one retained in so-called “toothless” pterosaurs like Pteranodon and Nyctosaurusthat have sharply tipped jaws.

Bigger Eggs?
A deeper pubis and pelvis in the BMNH specimen could have produced a larger egg. A stronger sternal complex and longer scapula could have made the “flapling” a more powerful flyer.

Soft Tissue Preservation
Despite a flipped mandible and poorly preserved feet, the “flapling” is otherwise well preserved and largely articulated. A soft tissue stain can be seen (overprinted in Fig. 1) that demonstrates a narrow chord at the elbow wing membrane construction.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

Updated September 30, 2016 with the addition of taxa to the LRT and new data on Protictis.

Where DO Bats Nest? The Question Returns.
A renowned (unnamed) professor interested in the origin of bats questioned my morphological nesting of bats with Ptilocercus and Nandinia (among living taxa) and Palaechthon (among fossil taxa). The professor sent me a pdf of Meredith et al. (2011), the most recent DNA tree to lump and split living mammals, as his best hypothesis on bat origins.

Figure 1. Bats and their sisters according to Meredith et al. 2011.

Mammal Diversification
Meredith et al. (2011) sought diversification patterns and times in mammals. They constructed a molecular supermatrix for mammalian families and analyzed these data with likelihood-based methods and relaxed molecular clocks. Their results came in traditionally, with Monotremata, Marsupialia and Placentalia at the base. The latter was divided into Xenartha + Afrotheria and all other placentals, which were divided into Laurasiatheria and Euarchontoglires.

DNA Results for Bats
Lots of bats were tested and they all lumped together in a single clade subdivided into three with fruit bats (Fig. 1 in orange) separating two microbat clades (in blue and green). Bats appeared as the unresolved sisters to Carnivora and Artiodactylia. Basal insectivores (not shown hre) nested as outgroup taxa to this super clade.

That’s an overly general nesting for bats that doesn’t provide much insight. On the other hand, I wasn’t surprised to see bats nesting so close to basal carnivores, like Nandinia and the vivverids, because the morphological results recovered the same relationship. I was surprised to bats nesting close to rhinos and camels. :-) Pangolins are indeed close to bats, so we agree here (Fig. 2).

DNA Results for Flying LemursThe base of the Euarchontoglires (Meredith et al. 2011, not shown in Fig. 1) included tree shrews and demopterans. I wasn’t surprised to see rabbits nesting close to Tupaia, the common tree shrew, because the morphological results recovered the same relationship. I also wasn’t surprised to see Ptilocercus, the pen-tailed tree shrew, nesting close to the flying lemurs, because the morphological results recovered the same relationship. Note these taxa didn’t nest with bats in the DNA study, but they did all nest at or near their unresolved common base.

Figure 2. Known bat ancestors to scale. Click to enlarge.

Morphological Results
The Meredith et al. (2011) results do not match the morphological evidence, which derives both bats and flying lemurs from a sister to Ptilocercus, a Paleocene pro primate and Chriacus, all close to basal carnivorans like Nandinia. Nandinia is a living carnivore that sometimes drops from trees and has an omniovorous diet. Chriacus was a long-legged tree-dwelling omnivore. Phylogenetic bracketing indicates that post-cranial characters were something like Chriacus and/or Ptilocercus. Ptilocercus is a flying lemur ancestor, but shares with bats several characters including flat ribs, a high floating scapula, wide cervicals, a rotating carpus and metatarsal + phalanx ratio similarities.

The question is…why don’t the DNA results more closely match the morphological results, and vice versa?

The DNA of modern tree shrews and bats, etc. is not the same as the DNA of Paleocene tree shrews and bats, etc.

The Meredith et al. (2011) evidence indicates that DNA results for large clades of mammals cannot resolve large clades. DNA and amino acid results do not agree with one another in the case of large reptile clades and the same is true in large mammal clades. DNA and amino acids apparently become more useful the more closely taxa are related. The resolution is very high, for instance, in human DNA, which is why it can be used in criminal investigations.

On the Other Hand
In fossil evidence you can point to a long list or suite of homologous morphologies, from tooth cusps to phalanx ratios. DNA results cannot provides these details. Morphology will always trump DNA, especially when bats nest with camels in DNA studies. DNA can only be verified with morphological evidence. DNA results can guide our efforts but the bottom line is morphology. The Meredith et al. (2011) study was unable to provide a specific sister taxon to bats. The morphological study provided Chriacus. When closer sisters are discovered, they will be reported.

Dermopterans and Bats
Flying lemurs nested close to bats and bat babys have short fingers like those of flying lemurs. Problem is: Ptilocercus, which comes between the two, has no extradermal membranes or webbed fingers and its limbs are not elongated. I have no answers for that other than both bats and flying lemurs are about 60 million years old and likely had a common long-limbed ancestor with extradermal membranes in a sister to Ptilocercus. Or bats and flying lemurs both developed extradermal membranes by convergence. Or Ptilocercus lost its ancestral long limbs and membranes.

Can we trust results?In science we don’t trust anything. Not DNA. Not morphology. Everything is tentative and provisional.

Eichstattisaurus and its sister, Ardeosaurus, were two small lizards found in Solnhofen limestones from the late Jurassic period, approximately 150 mya. Originally (Meyer 1860) and subsequently (Mateer 1982) these two were considered basal gekkotans, relatives of the living gecko, Gekko. Not much attention has been paid to either one. Both are typicall preserved complete and articulated, sometimes with some scalation and soft tissue preservation.

Figure 1. Eichstattisaurus and Ardeosaurus. Two Jurassic lizards in the lineage of snakes.

New Nesting
After entering the characters of both lizards, I was surprised to see that they nested not with Gekko, but with Adriosaurus, a hyper-elongated lizard with tiny limbs and a long neck, and Pachyrhachis, a basal snake with tiny hind limbs. This nesting, ancestral to snakes, has been largely overlooked by prior studies. I missed it too. The relationships is not obvious at first glance. I just finally got around to studying these two, fully expecting them to nest with Gekko.

Characters shared by members of this clade with Adriosaurus and Pachyrhachis include the orbit shape, the quadrate shape, supraoccipital fusion, converging temples, ectopterygoid shape, the absence of the retroarticular process, and the metatarsus configuration, among dozens of other traits that are shared with larger clades and by convergence with other reptilian clades.

The slender and elongated premaxillary ascending process was overlooked by Mateer (1982). If anyone has a palate view of either taxon, I’d like to see it.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

Kellner 2003
Dr. Alexander Kellner also nested Campylognathoides with Eudimorphodon but nested Rhamphorhynchus alone at the base of all pterodactyloid-grade pterosaurs. Dorygnathus, Scaphognathus and Sordes all nested at more basal positions.

The Present Tree
The present large tree, several times larger than any prior tree, and the first and only one to employ more than one specimen from several genera, nested the several species of Rhamphorhynchus following the several species of Campylognathoides and this clade was derived fromEudimorphodon cromptonellusandEudimorphodon ranzii.

No Consensus
It is apparent that no one here agrees with each other, but some share certain elements. Importantly no prior trees nested Rhamphorhynchus with Campylognathoides. This is likely due to the choice of which specimen was used in analysis. The variety within each genus is substantial and certain Rhamphorhynchus specimens do indeed converge with certain Dorygnathus specimens. The large study promoted here used several specimens in order to alleviate this problem. However, what we’re most interested in today is the Campylognathoides to Rhamphorhynchus transition.

Figure 1. The size reduction at the Campylognathoides to Rhamphorhynchus transition. From left to right: CM 11424, the Pittsburgh specimen of Campylognathoides, St/Ei 8209 Rhamphorhynchus intermedius, and the BMM specimen of Rhamphorhynchus.

Our Transitional Players
The most derived Campylognathoides is the Pittsburgh specimenCM 11424, specimen C3 in the Wild (1975 catalog) from the Early Jurassic. The most basal Rhamphorhynchus is R. intermedius(Koh 1937) , St/Ei 8209, No. 28 in the Wellnhofer 1975 catalog from the Late Jurassic. Not surprisingly, the latter looks like a smaller version of the former and had plenty of time to evolve from it. We know of no Campys in the Late Jurassic and no Rhamphs in the Early Jurassic.

A juvenile?
R. intermedius was considered a juvenile Rhamphorhychus by Bennett (1995), who used long bone measurements rather than a phylogenetic analysis. R. intermedius was larger than its phylogenetic successors, like R. longicaudus, but smaller than derived Rhamphorhynchus species, like R. longiceps. The phylogenetic size decrease between the specimens was due to serial precocious maturity and serial smaller egg size, as in several other pterosaur lineages.

Differences
Distinct from the C. liasicus, the skull of R. intermedius was relatively larger with a smaller naris and antorbital fenestra. Only one maxillary tooth was enlarged to fang status and like the premaxillary teeth, it was procumbent. The mandible was robust and convex dorsally. Several anterior dentary teeth also leaned anteriorly. The cervicals were slightly longer. The dorsal series was slightly shorter. The scapula and coracoid were not fused. This lack of fusion is not a sign of maturity, but follows phylogenetic lines. The deltopectoral crest was narrower. The ulna + radius was longer. The three distal wing phalanges were shorter and gracile. The prepubis perforation is expanded beyond the leading edge leaving an anterior process and a ventral process above and below the former perforation. The hind limbs were among the shortest among pterosaurs. The pedal digits were shorter than the metatarsals and digit V was longer than in Campylognathoides.

Size Reduction
In pterosaurs phylogenetic size reduction appears to mimic juvenile characters. But we already know that pterosaur hatchlings were nearly identical to adults. That means the phylogenetic changes precede that hatchling stage and move back into the embryonic stage. Smaller pterosaur adults matured more rapidly than larger pterosaur adults. Smaller pterosaur eggs were ready to hatch sooner than larger pterosaur eggs. These changes produced the smaller wings, tail and legs seen in R. intermedius.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

There are several skulls and fewer post-crania attributed to Youngina and Youngoides, originally by R. Broom (1914), but CE Gow (1975) and others (see below) also made contributions.

The big question is: are the skulls crushed into a variety of shapes? Or do the variety of shapes reflect important morphologies that separate the various specimens into various clades? If you have any Youngina/Youngoides skull photos, please send them!!

The other question is: do some specimens harbor an antorbital fenestra?

Here’s why I wonder:

Figure 1. Youngina BPI 375. Is this a nascent antorbital fenestra?

And Here’s Another One:

Figure 2. Youngina AMNH 5561. Is this a nascent antorbital fenestra?

At the Base of the Archosauriformes
These two Youngina specimens nest at the base of the Archosauriformes in the midst of several other younginiforms. Do those little skull breaks/indentations represent antorbital fenestra? Good question. The answer is, it really doesn’t matter in phylogenetic analysis because predecessors in the protorosauria do not have an antorbital fenestra and successors in the archosauriformes do. Not all Youngina had or have to have an antorbital fenestra. These things tend to come and go, especially when they first appear.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

ReferencesBroom R 1914. A new thecodont reptile. Proceedings of the Zoological Society of London, 1914:1072-1077.Gardner NM, Holliday CM and O’Keefe FR2010. The braincase of Youngina capensis(Reptilia, Diapsida): New insights from high-resolution CT scanning of the holotype. Paleonotologica Electronica 13(3):online PDFGow CE 1975. The morphology and relationships of Youngina capensis Broom and Prolacerta broomi Parrington. Palaeontologia Africana, 18:89-131.Olsen EC 1936. Notes on the skull of Youngina capensis Broom. Journal of Geology, 44 (4): 523-533.Reisz RR, Modesto SP and Scot DMT 2011. A new Early Permian reptile and its significance in early diapsid evolution. Proceedings of the Royal Society, London B
doi:10.1098/rspb.2011.0439